Individual diet and habitat specialisation are widespread in animal taxa and often related to levels of predation and competition. Mobile consumers such as predatory fish can stabilise lake food webs by ranging over a larger area than their prey, thereby switching between habitats. Although, this switching assumes that the predator has equal preference for the available prey, individual diet specialisation and morphological adaptations to different habitats could potentially prevent individuals from switching between habitats. In this study, we assessed the niche width and individual specialisation in Eurasian perch Perca fluviatilis in response to a shift in habitat use by manipulating the ability for this top predator to couple habitats. We ran an eight weeks pond experiment, to test the effect of habitat switching on diet and morphological specialisations. We show that habitat coupling influenced individual diet specialisation and niche use in expected directions where specialisation increased with decreasing habitat switching. In contrast to expectations, the morphological variation decreased with increasing diet specialisation. Our results expand on previous work and suggest that individual specialisation and niche width can impact the ability of mobile predators to couple habitats. Furthermore, it shows the importance of individual specialisations in relation to habitat coupling.

Trophic polymorphism has found to be common in many taxa and is a suggested mechanism of ecological speciation. To characterize the trophic linkages of specific morphotypes of organisms as well as a time-integrated niche use, several methods are available. In this study, we present data of multiple techniques to investigate the trophic divergence of Eurasian perch (Perca fluviatilis) that displays well-studied trophic polymorphism associated with littoral and pelagic habitats in lakes. We combined bulk stable isotope and fatty acid analyses on the muscle tissue of perch from three different lakes in Sweden with analyses of stomach content. By comparing the three methods, we aimed at providing a broad and highly resolved picture on the trophic divergence in freshwater fish. The degree in morphological divergence varied between perch caught in the three different lakes. Generally, perch caught in the pelagic zone were more streamlined compared to the ones caught in the littoral zone that had a deeper body, as shown by geometric morphometrics. The three diet assessment methods revealed different levels of information. Data on stomach content showed some preferences for specific dietary items in littoral and pelagic perch, but general trophic specialization could not be concluded due to the small sample size. Analyses of delta C-13 and delta N-15, however, confirmed these results as a long-term pattern connected to specific habitat use in two of the three lakes. Fatty acid signatures of perch reflected partly those of the prey items of the specific habitats. Although the proportions of the essential fatty acid 22:6n-3 were lower in littoral resources, the proportions in littoral fish were similar to the ones caught in the pelagic zone. We concluded that although a fundamental contribution from littoral resources exists in littoral phenotypes, a minor reliance on pelagic prey items is obviously needed to provide essential compounds. Thus, by combining the methods to characterize direct resource use (i.e., stomach analyses) with others that utilize trophic biomarkers (i.e., analyses of stable isotopes and fatty acids), we were able to illustrate the degree of variation in trophic divergence of perch but also shed some light on potential trade-offs that are related to resource specialization in freshwater fish.

Increasing input of terrestrial dissolved organic carbon (DOC) has been identified as a widespread environmental phenomenon in many aquatic ecosystems. Terrestrial DOC influences basal trophic levels: it can subsidize pelagic bacterial production and impede benthic primary production via light attenuation. However, little is known about the impacts of elevated DOC concentrations on higher trophic levels, especially on top consumers. Here, we used Eurasian perch (Perca fluviatilis) to investigate the effects of increasing DOC concentrations on top predator populations. We applied stable isotope analysis and geometric morphometrics to estimate long-term resource and habitat utilization of perch. Habitat coupling, the ability to exploit littoral and pelagic resources, strongly decreased with increasing DOC concentrations due to a shift toward feeding predominantly on pelagic resources. Simultaneously, resource use and body morphology became increasingly alike for littoral and pelagic perch populations with increasing DOC, suggesting more intense competition in lakes with high DOC. Eye size of perch increased with increasing DOC concentrations, likely as a result of deteriorating visual conditions, suggesting a sensory response to environmental change. Increasing input of DOC to aquatic ecosystems is a common result of environmental change and might affect top predator populations in multiple and complex ways.

Individuals are constantly in competition with one another and, on both ecological and evolutionary timescales, processes act to reduce this competition and promote the gain of fitness advantages via diversification. Here we have investigated the genetic (AFLP) and morphological (geometric morphometrics) aspects of the littoral–pelagic axis, a commonly observed resource polymorphism in freshwater fishes of postglacial lakes. We found a large degree of variation in the genetic and morphological divergence between littoral and pelagic perch and roach across Swedish lakes. Although there was evidence of assortative mating (elevated kinship values) in both species, we could not find any significant coupling of morphology and genetic divergence. Instead, there was evidence that the extent of resource polymorphism may be largely caused by phenotypic plasticity. These results suggest that assortative mating, which can lead to genetically determined adaptive divergence, does occur in these species, particularly perch, but not according to genetically fixed morphological traits. The behavioural mechanisms facilitating associative mating need to be investigated to explore the interaction between phenotypic plasticity and adaptive genetic divergence and their roles in diversification.

Among-individual diet variation is common in natural populations and may occur at any trophic level within a food web. Yet, little is known about its variation among trophic levels and how such variation could affect phenotypic divergence within populations. In this study we investigate the relationships between trophic position (the population’s range and average) and among-individual diet variation. We test for diet variation among individuals and across size classes of Eurasian perch (Perca fluviatilis), a widespread predatory freshwater fish that undergoes ontogenetic niche shifts. Second, we investigate among-individual diet variation within fish and invertebrate populations in two different lake communities using stable isotopes. Third, we test potential evolutionary implications of population trophic position by assessing the relationship between the proportion of piscivorous perch (populations of higher trophic position) and the degree of phenotypic divergence between littoral and pelagic perch sub-populations. We show that among-individual diet variation is highest at intermediate trophic positions, and that this high degree of among-individual variation likely causes an increase in the range of trophic positions among individuals. We also found that phenotypic divergence was negatively related to trophic position in a population. This study thus shows that trophic position is related to and may be important for among-individual diet variation as well as to phenotypic divergence within populations.

A positive relationship between occupancy and average local abundance of species is found in a variety of taxa, yet the mechanisms driving this association between abundance and occupancy are still enigmatic. Here we show that freshwater fishes exhibit a positive abundance-occupancy relationship across 125 Swedish lakes. For a subset of 9 species from 11 lakes, we estimated species-specific diet breadth from stable isotopes, within-lake habitat breadth from catch data for littoral and pelagic nets, adaptive potential from genetic diversity, abiotic niche position, and dispersal capacity. Average local abundance was mainly positively associated with both within-lake habitat and diet breadth, that is, species with larger intraspecific variation in niche space had higher abundances. No measure was a good predictor of occupancy, indicating that occupancy may be more directly related to abundance or abiotic conditions than to niche breadth per se. This study suggests a link between intraspecific niche variation and a positive abundance-occupancy relationship and implies that management of freshwater fish communities, whether to conserve threatened or control invasive species, should initially be aimed at niche processes.

Habitat protection is a strategy often proposed in fisheries management to help maintain viable populations of exploited species. Yet, quantifying the importance of habitat availability for population sizes is difficult, as the precise distribution of essential habitats is poorly known. To quantify the contribution from coastal nursery habitats to exploited fish population sizes, we related adult density to the amount of nursery habitat available for 12 populations of the two dominant predatory fish species in a 40 000-km² archipelago area of the Baltic Sea. Habitat distribution was mapped using three conceptually different techniques, Maxent, generalized additive models, and random forest, using spawning and 0-group point samples. Adult densities were estimated from gillnet surveys. Regressions demonstrated no evident effect from fishing, whereas habitat availability had a positive effect, explaining almost half of the variation in population sizes of both species. This result shows that a substantial proportion of the potential production of adult fish can be estimated by mapping essential nursery habitats distribution. Responses were non-linear, indicating that habitat protection has largest effects where there is little available habitat. By demonstrating the importance of habitat limitation of two exploited fish species, we provide quantitative support to the benefits of habitat protection for fisheries.

While phenotypic responses to direct species interactions are well studied, we know little about the consequences of indirect interactions for phenotypic divergence.In this study we used lakes with and without the zebra mussel to investigate effects ofindirect trophic interactions on phenotypic divergence between littoral and pelagic perch. We found a greater phenotypic divergence between littoral and pelagic individuals inlakes with zebra mussels and propose a mussel-mediated increase in pelagic and benthic resource availability as a major factor underlying this divergence. Lakes withzebra mussels contained higher densities of large plankton taxa and large invertebrates. We suggest that this augmented resource availability improved perch foraging opportunities in both the littoral and pelagic zones. Perch in both habitats could hence express a more specialized foraging morphology, leading to an increased divergence of perch forms in lakes with zebra mussels. As perch do not prey on mussels directly, we conclude that the increased divergence results from indirect interactions with the mussels. Our results hence suggest that species at lower food web levels can indirectlyaffect phenotypic divergence in species at the top of the food chain.

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C-4 plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (delta C-13). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.